1. Department of Biological and Environmental Science; University of Jyvaskyla; P.O. Box 35 40014 Jyvaskyla Finland
2. Ecology, Evolution & Genetics, Research School of Biology; Australian National University; Canberra ACT 0200 Australia
3. Institute of Evolutionary Biology and Environmental Studies; University of Zurich; Winterthurerstrasse 190 CH-8057 Zurich Switzerland